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Choosing

Lynch
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Choosing HEALTH
help you get back on track. Health Coaching Activities also have hints
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APRIL LYNCH
BARRY ELMORE

THIRD EDITION
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CHOOSING
HEALTH THIRD EDITION

April Lynch, M.A.


Karen Vail-Smith, M.S., M.P.A.
EAST CAROLINA UNIVERSITY

Jerome Kotecki, H.S.D.


BALL STATE UNIVERSITY

With contributions by
Laura Bonazzoli

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Credits and acknowledgments borrowed from other sources and reproduced, with permission, in
this textbook appear on the appropriate page within the text or on page CR-1.
Copyright © 2018, 2015, 2012 by Pearson Education, Inc. Published by Pearson Education, Inc.
All rights reserved. Manufactured in the United States of America. This publication is protected
by Copyright and permission should be obtained from the publisher prior to any prohibited
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aware of a trademark claim, the designations have been printed in initial caps or all caps.
pearson, always learning, MasteringHealth®, are exclusive trademarks, in the United States and/or
other countries, of Pearson Education, Inc. or its affiliates.

Library of Congress Cataloging-in-Publication Data on file.

ISBN 10: 0-134-49367-2; ISBN 13: 978-0-134-49367-1 (Student edition)


ISBN 10: 0-134-63624-4; ISBN 13: 978-0-134-63624-5 (Instructor’s Review Copy)
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1 16

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“ This book is dedicated to my husband,
Colin, daughter, Ava, and son, Van. In the
ever-changing love and laughter project
that is our family, I’m inspired to reach for
better choices, every single day.”
—April Lynch

“ IWilliam,
dedicate this book to my loving family,
Alex, Mary-Brett, and Zan. I thank
my colleagues, who always provide support
and good counsel, and my students, who
teach me every day.”
—Karen
Vail-Smith

“ This book is dedicated to my friends and


family for their continued support and love.
They allowed me the time and energy to
focus my passion and write this book.”
—Jerome Kotecki

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ABOUT THE AUTHORS

April Lynch, M.A.


April Lynch is an award-winning author and journalist who specializes in health, the medical
and biological sciences, and human genetics. During her tenure with the San Jose Mercury
News, the leading newspaper of Silicon Valley, she served as the science and health editor,
focusing the paper’s coverage on personal health and scientific developments in the field of
disease prevention. She has also worked as a writer and editor for the San Francisco Chronicle.
April has written numerous articles on personal health, medical and scientific advances,
consumer issues such as health insurance, and the ways that scientific breakthroughs are
redefining our understanding of health. She has been a frequent contributor to leading
university textbooks covering applied biology, nutrition, and environmental health and science.
Along with Choosing Health, April has co-authored Health: Making Choices for Life, an
innovative personal health textbook for instructors who desire a more detailed, in-depth book
for students majoring in health-related subjects. Together with a leading genetic counselor, April
is also the co-author of The Genome Book, a hands-on guide to using genetic information in
personal health decisions. Her work has won numerous awards from organizations such as the
Society of Professional Journalists, the California Newspaper Publishers Association, and the
Associated Press. Her current interests include a focus on how people receive and interact with
health information online as well as how complex scientific and medical information is best
shared compellingly and effectively in digital media. She lives in the San Francisco Bay Area with
her husband and children.

Karen Vail-Smith, M.S., M.P.A.


East Carolina University
Karen Vail-Smith received a B.S. from The University of North Carolina at Chapel Hill and an
M.S. and an M.P.A. from East Carolina University. She has been a faculty member in East
Carolina University’s Department of Health Education and Promotion for 27 years and has
taught more than 10,000 college students how to lead healthier lives. She specializes in
personal health, alcohol and other drugs, and human sexuality. She has received numerous
teaching awards, including the prestigious UNC Board of Governor’s Distinguished Professor
for Teaching award. During her career she has published more than 30 articles in health
professional journals, produced 50 nationally distributed health promotion videos, and
contributed to several personal health textbooks.

Jerome Kotecki, H.S.D.


Ball State University
Jerome E. Kotecki is a professor of health science in the Department of Nutrition and Health
Science, located within the College of Health, at Ball State University. Jerome earned his
doctorate in health education and his master’s degree in exercise science from Indiana
University. He has published more than 40 scientific research papers on the prevention, arrest,
and reversal of the most common chronic diseases facing Americans today. Jerome has
authored and co-authored multiple textbooks on the importance of healthy lifestyle habits
to enhance the multidimensional components of human health and prevent cardiovascular
disease, diabetes, cancer, and other chronic conditions. Jerome has extensive experience
in health promotion, with particular focus on physical activity and health. An experienced
teacher and researcher, he is devoted to helping students adopt and maintain healthy lifestyles.
Jerome has been recognized for his contributions to the scholarship of teaching and learning
by his department, college, and university. He is an avid fitness participant and enjoys cycling,
resistance training, running, mountain biking, hiking, swimming, and yoga.

iv

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ABOUT THE CONTRIBUTOR

Laura Bonazzoli
Laura Bonazzoli has been writing and editing in the health sciences for over 20 years. Her early
work in human anatomy and physiology, chemistry, and other core sciences laid the foundation
for writing projects in nursing, pathology, nutrition, complementary and alternative medicine,
and personal health. Her commitment as a writer is to help her readers appreciate the power
of small choices to improve their health and the health of their communities. In her free time,
Laura and her daughter enjoy exploring the gardens, byways, and beaches of mid-coast Maine.

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BRIEF CONTENTS

1 HEALTH IN THE 21ST CENTURY 1

2 PSYCHOLOGICAL HEALTH 25

3 STRESS MANAGEMENT 51

4 SLEEP, YOUR BODY, AND YOUR MIND 74

5 NUTRITION 96

6 PHYSICAL ACTIVITY FOR FITNESS AND HEALTH 126

7 BODY IMAGE, BODY WEIGHT 154

8 ADDICTIONS AND DRUG USE 181

9 ALCOHOL AND TOBACCO USE AND ABUSE 205

10 SOCIAL RELATIONSHIPS AND COMMUNICATION 234

11 SEXUALITY, CONTRACEPTION, AND REPRODUCTIVE CHOICES 255

12 PREVENTING INFECTIOUS DISEASES AND SEXUALLY


TRANSMITTED INFECTIONS 291

13 DIABETES, CARDIOVASCULAR DISEASE, AND CANCER 318

14 CONSUMER HEALTH 352

15 PERSONAL SAFETY AND INJURY PREVENTION 376

Additional Electronic Chapters


Access these chapters online through

16 YOUR ENVIRONMENT, YOUR HEALTH 400

17 AGING WELL 430

vii

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CONTENTS

Preface xxiii | Acknowledgments xxix

CHAPTER 1 Personal Choices 15


HEALTH IN THE 21ST CENTURY 1 Campus and Community Advocacy 18
Choosing to Change Worksheet 21
Chapter Summary 22
Get Connected 23
Test Your Knowledge 23
What Do You Think? 24

CHAPTER 2
PSYCHOLOGICAL HEALTH 25

What Is Health? 2
Health Versus Disease 2
Health Versus Wellness 3
Dimensions of Health 3
Current Health Challenges 4
Health Across America 4
Health on America’s Campuses 7
Health Around the World 7
Determinants of Health 8
Biology and Genetics 9 What Is Psychological Health? 26
Individual Behaviors 9 Components of Psychological Health 26
Social Determinants 10 Facets of Psychological Health 26
Physical Determinants 10 Maslow’s Hierarchy of Needs 28
Health Services 11 The Role of Emotional Intelligence 28
Policy-Making 11 The Value of Optimism 29

What Factors Influence Behavior Factors Affecting Psychological Health 29


Change? 13 Family History 29
Predisposing Factors 13 Social Support 29
Enabling Factors 13 The Role of Spiritual Health 29
Reinforcing Factors 13 Common Psychological Challenges 31
How Does Behavior Change Occur? 14 Shyness 31
The Transtheoretical Model 14 Loneliness 32
The Health Belief Model 14 Anger 32
Ecological Models 14
Mental Disorders in the United States: An
Change Yourself, Change Your World 15 Overview 33

viii

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Prevalence of Mental Disorders 33 What Is Stress? 52
Diagnosis and Treatment of Mental Disorders 33
The Body’s Stress Response 52
Mood Disorders 34 Alarm Stage: The Fight-or-Flight
Depressive Disorders 34 Response 53
Bipolar Disorder 36 Resistance Stage 54
Anxiety Disorders 37 Exhaustion Stage and Allostatic Overload 54
Generalized Anxiety Disorder (GAD) 37 Health Effects of Chronic
Panic Attacks and Panic Disorder 37 Stress 54
Social Anxiety Disorder 37 Effects on the Cardiovascular System 55
Phobias 37 Effects on Digestion, Glucose Regulation, and
Treating Anxiety Disorders 38 Body Weight 55
Other Disorders 38 Effects on the Immune System 56
Obsessive-Compulsive Disorder (OCD) 38 Stress and Headaches 56
Post-Traumatic Stress Disorder (PTSD) 38 Effects on Sleep 56
Attention Disorders 39 Effects on Relationships and Sexual
Schizophrenia 40 Functioning 57
Effects on Mind and Mental Health 57
Self-Injury and Suicide 40
Non-suicidal Self-Injury 40 What Influences the Stress
Suicide 41 Response? 57
The Role of Personality Types 57
Getting Help for a Psychological Problem 42
The Role of Personality Traits 58
Options on Campus 42
The Role of Biology 58
Clinical Options 42
What Are College Students’ Common
Change Yourself, Change Your World 45
Stressors? 59
Taking Care of Yourself 45
Academic Pressure 59
Helping Others 45
Financial Stressors 59
Choosing to Change Worksheet 46 Job-Related Stressors 59
Chapter Summary 48 Social Stressors 59
Get Connected 49 Minor Hassles and Major Life
Test Your Knowledge 49 Changes 61
What Do You Think? 50 Environmental Stressors 62
Internal Stressors 62
Resources for Managing Stress 62
CHAPTER 3 Social Support 62
STRESS MANAGEMENT 51 Help on Campus 62
Medical Options 64

Change Yourself, Change Your


World 64
Manage Your Time Effectively 64
Improve Your Test-Taking Skills 65
Live a Healthier Lifestyle 65
Relieve Your Tensions 66
Change Your Thinking 67
Create a Personalized Stress
Management Plan 68
Choosing to Change Worksheet 69
Chapter Summary 71
Get Connected 72
Test Your Knowledge 72
What Do You Think? 73

CONTENTS ix

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CHAPTER 4 CHAPTER 5
SLEEP, YOUR BODY, AND NUTRITION 96
YOUR MIND 74

What Are Nutrients? 97


Energy and Calories 97
What Is Sleep? 75
Carbohydrates 98
Regions and Rhythms of Sleep 75
Fats 100
Stages of Sleep 76
Proteins 102
Cycles of Sleep 77
Vitamins 103
Sleep: How Much and Why It Minerals 104
Matters 77 Water 104
Research on Short and Long Sleep 78 What About Functional Foods and Dietary
Short Sleep: The American Way? 79 Supplements? 106
Why Is Ample Sleep Important? 79
Phytochemicals 107
What Factors Influence Sleep? 81 Probiotics and Prebiotics 108
Biology and Genetics 81 Dietary Supplements 108
Individual Behaviors 81 What Tools Can Help You Eat Right? 110
Factors in the Environment 83
Dietary Reference Intakes (DRIs) 110
Sleep Disorders 83 Food Labels 111
Insomnia 83 Dietary Guidelines for Americans 113
Snoring 83 MyPlate 113
Sleep Apnea 84 Is Our Food Supply Safe? 116
Narcolepsy 85
Foodborne Illness 116
Parasomnias 85
Food Allergies and Intolerances 117
Getting Help for a Sleep Disorder 86 Food Residues 118
Campus and Community Support 86 Change Yourself, Change Your World 118
Clinical Diagnosis and Treatment 86
Think Smart When Making
Complementary and Alternative Choices 120
Therapies 88
Eat Smart When Eating Out 120
Change Yourself, Change Your World 88 Shop Smart When Money’s Tight 120
Personal Choices 88 Campus Advocacy 121

Choosing to Change Worksheet 91 Choosing to Change Worksheet 122


Chapter Summary 93 Chapter Summary 123
Get Connected 94 Get Connected 124
Test Your Knowledge 94 Test Your Knowledge 124
What Do You Think? 95 What Do You Think? 125

x CONTENTS

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CHAPTER 6 Exercise Safe, Exercise Smart 145
Get Medical Clearance and
PHYSICAL ACTIVITY FOR FITNESS Be Prepared 145
Warm Up and Cool Down 145
AND HEALTH 126 Get Training 145
Eat Right 145
Stay Hydrated 145
Prepare for Hot or Cold Weather 146
Start Slowly and Watch Out for Red Flags 146
Care for Injuries 147
Be Wary of Performance-Enhancing Drugs 148

Change Yourself, Change Your World 148


Personal Choices 148
Choosing to Change Worksheet 151
Chapter Summary 152
Get Connected 152
Test Your Knowledge 152
What Do You Think? 153

What Is Physical Fitness? 127


CHAPTER 7
Cardiorespiratory Fitness 127 BODY IMAGE, BODY WEIGHT 154
Muscular Strength 127
Muscular Endurance 128
Flexibility 128
Body Composition 128
What Are the Benefits of Physical
Activity? 129
Stronger Heart and Lungs 129
Management and Prevention of Type 2 Diabetes 130
Reduced Risk of Some Cancers 130
Increased Immune Function 130
Stronger Bones 130
Reduced Risk of Injury 130
Healthful Weight Management 130
Benefits to Psychological Health, Stress Management,
and Sleep 130
Principles of Fitness Training 131
Overload 131 Body Image and Body Weight 155
Specificity 131 Many Factors Influence Body Image 155
Reversibility 131 Defining a Healthful Body Weight 156
Individuality 132
Diminishing Returns 132 Alarming Trends in Body Weight 158
Weight Trends in the United States 158
What Types of Physical Activity Should You
Weight Trends Around the World 158
Consider? 132
Weight Trends on Campus 159
Aerobic Exercise 132
Exercise for Muscular Strength and Endurance 135 Risks and Costs of Obesity 159
Exercises for Improving Flexibility 136 Health Risks 159
How Much Physical Activity Do You Financial Burden of Obesity 160
Need? 142 Factors that Contribute to Weight Gain 161
Guidelines for Health Maintenance 142 Biology and Genetics 161
Avoid Sustained Sitting 144 Individual Behaviors 163
Increase Your Level of Activity 144 Social Factors 163

CONTENTS xi

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Physical Factors 164 Compulsive Spending 183
Public Policy 164 Technology Addiction 183
Reducing Excess Body Weight 164 Patterns of Drug Use 184
Can Dieting Work 164
Comparing Diets 166 How the Body Responds to Drugs 185
Diet “Aids”? 167 Drug Misuse and Abuse 185
Clinical Options for Obesity 168 Initial Effects on the Brain 185
The Importance of Physical Activity 169 Effects of Chronic Use 185
What If You Want To Gain Weight? 169 How Drugs Leave the Body 187
How Do You Maintain a Healthful Commonly Abused Drugs 187
Weight? 170 Prescription and Over-the-Counter
Change Yourself, Change Your World 170 Medications 187
Campus Resources for Maintaining a Healthy Heroin 188
Weight 171 Marijuana 190
Stimulants 191
Body Image and Eating Hallucinogens 193
Disorders 171 Club Drugs 193
Body Image Disorders 171 Inhalants 194
Eating Disorders 172 Depressants 194
Other Unhealthful Eating Behaviors 174
Getting Help for a Body Image or Eating Preventing Drug Abuse 195
Disorder 175 Getting Help for a Drug Problem 195
Personal Choices: Develop a More Positive Body
Campus and Community-Based Options 197
Image 176
Clinical Options 197
Choosing to Change Worksheet 176
Change Yourself, Change Your World 198
Chapter Summary 178
Personal Choices 198
Get Connected 179
Helping a Friend 199
Test Your Knowledge 179
What Do You Think? 180 Choosing to Change Worksheet 200
Chapter Summary 202
Get Connected 203
CHAPTER 8 Test Your Knowledge 203
What Do You Think? 204
ADDICTIONS AND DRUG USE 181
CHAPTER 9
ALCOHOL AND TOBACCO USE
AND ABUSE 205

An Overview of Addiction 182


What Is Addiction? 182
What Are Behavioral Addictions? 183
Pathological Gambling 183
Hypersexual Disorder 183

xii CONTENTS

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Alcohol Use in the United the States 206
Alcohol Use and Binge Drinking on Campus 206
CHAPTER 10
The Makeup of Alcohol 208 SOCIAL RELATIONSHIPS AND
How the Body Absorbs and Metabolizes COMMUNICATION 234
Alcohol 209
Blood Alcohol Concentration 209
The Effects of Alcohol on the Body 211
Intoxication 211
Immediate Effects of Alcohol on
the Body 211
Long-Term Effects of Alcohol on the Body 212
Alcohol and Pregnancy 213
Health Benefits of Alcohol? 214
The Effects of Alcohol on Behavior 214
Drinking and Driving 214
Alcohol and Sexual Activity 215
Alcohol and Other Problems 216
Alcohol Abuse 216
Alcoholism 216
Alcohol Abuse: Treatment and Communication in Relationships 235
Prevention 217 Communicating Feelings 235
Treatment Options 218 Being a Good Listener 236
Dealing with Relapse 218 Resolving Conflicts 236
Prevention Strategies 218 Gender Roles and Communication 237
Smoking in the United States 220 Developing Relationships 238
Smoking on Campus 220 Self-Perception 238
What’s in a Cigarette? 221 Early Relationships 238
Gender Roles 238
Effects of Smoking on Health 221
Short-Term Health Effects 221 Friendships 239
Long-Term Health Effects 223 Maintaining Old Friendships 239
Smoking and Pregnancy 223
Intimate Relationships 240
Secondhand Smoke 224
Sternberg’s Triangular Theory of Love 240
Other Forms of Tobacco 225 What Causes Attraction? 242
Cigars 225 Dating 242
Clove Cigarettes 225 Same-Sex Relationships 243
Bidis 225 Healthy Relationships 243
Hookahs 226 Dysfunctional Relationships 244
Smokeless (“Spit”) Tobacco 226 When Relationships End 245
Electronic Cigarettes 226
Committed Relationships 245
Getting Help to Quit Cohabitation 246
Smoking 227 Marriage 246
Treatment Options 227 Domestic Partnerships 248
Dealing with Relapse 228 Staying Single 248

Change Yourself, Change Your World 228 Starting a Family 248


Personal Choices 228 Choosing to Have Children 249
Campus Advocacy 229 Stepfamilies 249
Single Parenthood 249
Choosing to Change Worksheet 230
Characteristics of Happy Families 250
Chapter Summary 231
Get Connected 232 Change Yourself, Change Your World 250
Test Your Knowledge 232 Personal Choices 250
What Do You Think? 233 Campus Advocacy 250

CONTENTS xiii

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Choosing to Change Worksheet 251 Methods of Abortion 278
Chapter Summary 253 Physical and Psychological Complications of
Get Connected 253 Abortion 279
Test Your Knowledge 254 Legal Status of Abortion 279
What Do You Think? 254
Pregnancy and Childbirth 280
Pregnancy 280

11
Childbirth 284
CHAPTER
Infertility 285
SEXUALITY, CONTRACEPTION, Causes of Infertility 285
Options for Infertile Couples 286
AND REPRODUCTIVE Change Yourself, Change Your World 286
CHOICES 255 Personal Choices 286
Campus Advocacy 287
Choosing to Change Worksheet 287
Chapter Summary 288
Get Connected 289
Test Your Knowledge 290
What Do You Think? 290

CHAPTER 12
PREVENTING INFECTIOUS
DISEASES AND SEXUALLY
TRANSMITTED INFECTIONS 291

Sexual Anatomy and Health 256


Female Sexual Anatomy and Sexual Health 256
Male Sexual Anatomy and Sexual Health 258
The Menstrual Cycle 259
Phases of the Menstrual Cycle 260
Disorders Associated with the Menstrual Cycle 261
The Sexual Response Cycle 262
Sexual Dysfunctions 262
Sexual Behavior 264
Abstinence and Celibacy 264
Sexual Intercourse 264
Oral Sex 264
Communicating About Sex 265
Non-Intercourse Sexual Activity 265 How Are Infections Spread? 292

Sexual Orientation and Gender Identity 267 Protecting Against Infections 293
Heterosexuality 267 The Body’s First Line of Defense 293
Homosexuality 267 The Body’s Immune Response 293
Bisexuality 268 Immunization 294
Gender Identity 268 Immune Disorders 296
Infectious Diseases 297
Conception and Contraception 268
Viral Infections 297
Conception 269
Bacterial Infections 299
Contraceptive Options 269
Fungal Infections 303
Which Method Is the Best? 276
Protozoan Infections 303
Abortion 277 Parasitic Worm Infections 304

xiv CONTENTS

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The number of joints of the antennae increases during the
development; it is effected by growth of the third joint and
subsequent division thereof; hence the joints immediately beyond
the second are younger than the others, and are usually shorter and
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when we recollect that these creatures live in communities
concealed from observation, and that an isolated individual cannot
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Natural History.—The progress of knowledge as to Termites has


shown that profound differences exist in the economy of different
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from one species. We will therefore briefly sketch the economy, so
far as it has been ascertained, in three species, viz. Calotermes
flavicollis, Termes lucifugus, and T. bellicosus.

Fig. 230.—Some individuals of Calotermes flavicollis: A, nymph with


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individual. (After Grassi.)

Calotermes flavicollis inhabits the neighbourhood of the


Mediterranean Sea; it is a representative of a large series of species
in which the peculiarities of Termite life are exhibited in a
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members of the community, viz. soldiers, and the young and
adolescent. The habits of this species have recently been studied in
detail in Sicily by Grassi and Sandias.[289] The Insects dwell in the
branches and stems of decaying or even dead trees, where they
nourish themselves on those parts of the wood in which the process
of decay is not far advanced; they live in the interior of the stems, so
that frequently no sign of them can be seen outside, even though
they may be heard at work by applying the ear to a branch. They
form no special habitation, the interior of the branch being sufficient
protection, but they excavate or increase the natural cavities to suit
their purposes. It is said that they line the galleries with proctodaeal
cement; this is doubtful, but they form barricades and partitions
where necessary, by cementing together the proctodaeal products
with matter from the salivary glands or regurgitated from the anterior
parts of the alimentary canal. The numbers of a community only
increase slowly and remain always small; rarely do they reach 1000,
and usually remain very much below this. The king and queen move
about, and their family increases but slowly. After fifteen months of
their union they may be surrounded by fifteen or twenty young; in
another twelve months the number may have increased to fifty, and
by the time it has reached some five hundred or upwards the
increase ceases. This is due to the fact that the fertility of the queen
is at first progressive, but ceases to be so. A queen three or four
years old produces at the time of maximum production four to six
eggs a day. When the community is small—during its first two years
—the winged individuals that depart from it are about eight or ten
annually, but the numbers of the swarm augment with the increase of
the population. The growth of the individuals is slow; it appears that
more than a year elapses between the hatching of the egg and the
development of the winged Insect. The soldier may complete its
development in less than a year; the duration of its life is not known;
that of the kings and queens must be four or five years, probably
more. After the winged Insects leave the colony they associate
themselves in pairs, each of which should, if all goes well, start a
new colony.

The economy of Termes lucifugus, the only European Termite


besides Calotermes flavicollis, has been studied by several
observers, the most important of whom are Lespès[290] and Grassi
and Sandias. This species is much more advanced in social life than
Calotermes is, and possesses both workers and soldiers (Fig. 231,
2, 3); the individuals are much smaller than those of Calotermes.
Burrows are made in wood of various kinds, furniture being
sometimes attacked. Besides making excavations this species builds
galleries, so that it can move from one object to another without
being exposed; it being a rule—subject to certain exceptions—that
Termites will not expose themselves in the outer air. This is probably
due not only to the necessity for protection against enemies, but also
to the fact that they cannot bear a dry atmosphere; if exposed to a
drying air they speedily succumb. Occasionally specimens may be
seen at large; Grassi considers these to be merely explorers. Owing
to the extent of the colonies it is difficult to estimate with accuracy
the number of individuals composing a community, but it is doubtless
a great many thousands. Grassi finds the economy of this species in
Sicily to be different from anything that has been recorded as
occurring in other species; there is never a true royal pair. He says
that during a period of six years he has examined thousands of nests
without ever finding such a pair. In place thereof there are a
considerable number of complementary queens—that is, females
that have not gone through the full development to perfect Insects,
but have been arrested in various stages of development. In Fig.
231, Nos. 4 and 5 show two of these abnormal royalties; No. 4 is
comparatively juvenile in form, while No. 5 is an individual that has
been substituted in an orphaned nest, and is nearer to the natural
condition of perfect development. We have no information as to
whether any development goes on in these individuals after the state
of royalty is assumed, or whether the differences between these
neoteinic queens are due to the state of development they may
happen to be in when adopted as royalties. Kings are not usually
present in these Sicilian nests; twice only has Grassi found a king,
but he thinks that had he been able to search in the months of
August and September he would then have found kings. It would
appear therefore that the complementary kings die, or are killed after
they have fertilised the females. Parthenogenesis is not thought to
occur, as Grassi has found the spermathecae of the complementary
queens to contain spermatozoa.
Fig. 231.—Some of the forms of Termes lucifugus. 1, Young larva; 2, adult
worker; 3, soldier; 4, young complementary queen; 5, older substitution
queen; 6, perfect winged Insect. (After Grassi.)

The period of development apparently occupies from eighteen to twenty-


three months. At intervals swarms of a great number of winged
individuals leave the nest, and are usually promptly eaten up by various
animals. After swarming, the wings are thrown off, and sometimes two
specimens or three may be seen running off together; this has been
supposed to be preliminary to pairing, but Grassi says this is not the case,
but that the object is to obtain their favourite food, as we shall mention
subsequently.

Although these are the usual habits of Termes lucifugus at present in


Sicily, it must not be concluded that they are invariable; we have in fact
evidence to the contrary. Grassi has himself been able to procure in
confinement a colony—or rather the commencement of one—
accompanied by a true royal pair; while Perris has recorded[291] that in
the Landes he frequently found a royal pair of T. lucifugus under chips;
they were accompanied in nearly every case by a few eggs. And
Professor Perez has recently placed a winged pair of this species in a box
with some wood, with the result that after some months a young colony
has been founded. It appears probable therefore that this species at times
establishes new colonies by means of royal pairs derived from winged
individuals, but after their establishment maintains such colonies as long
as possible by means of complementary queens. It is far from improbable
that distinctions as to the use of true and complementary royalties may be
to some extent due to climatic conditions. In some localities T. lucifugus
has multiplied to such an extent as to be very injurious, while in others
where it is found it has never been known to do so.

The Termitidae of Africa are the most remarkable that have yet been
discovered, and it is probably on that continent that the results of the
Termitid economy have reached their climax. Our knowledge of the
Termites of tropical Africa is chiefly due to Smeathman, who has
described the habits of several species, among them T. bellicosus. It is
more than a century since Smeathman travelled in Africa and read an
account of the Termites to the Royal Society.[292] His information was the
first of any importance about Termitidae that was given to the world; it is,
as may be well understood, deficient in many details, but is nevertheless
of great value. Though his statements have been doubted they are
truthful, and have been confirmed by Savage.[293]

Fig. 232.—Royal cell of Termes bellicosus, partially broken open to show


the queen and her attendants. (After Smeathman.) B, Antenna of the
queen; b, b, line of entrances to the cell; A, A, an entrance, in this line,
closed by the Termites. × ⅞.

T. bellicosus forms buildings comparable to human dwellings; some of


them being twenty feet in height and of great solidity. In some parts of
West Africa these nests were, in Smeathman's time, so numerous that
they had the appearance of villages. Each nest was the centre of a
community of countless numbers of individuals; subterranean passages
extended from them in various directions. The variety of forms in one of
these communities has not been well ascertained, but it would seem that
the division of labour is carried to a great extent. The soldiers are fifteen
times the size of the workers. The community is dependent on one royal
couple. It is the opinion of the natives that if that couple perish so also
does the community; and if this be correct we may conclude that this
species has not a perfect system of replacing royal couples. The queen
attains an almost incredible size and fertility. Smeathman noticed the
great and gradual growth of the abdomen, and says it enlarges "to such
an enormous size that an old queen will have it increased so as to be
fifteen hundred or two thousand times the bulk of the rest of her body, and
twenty or thirty thousand times the bulk of a labourer, as I have found by
carefully weighing and computing the different states." He also describes
the rate at which the eggs are produced, saying that there is a constant
peristaltic movement[294] of the abdomen, "so that one part or other
alternately is rising and sinking in perpetual succession, and the matrix
seems never at rest, but is always protruding eggs to the amount (as I
have frequently counted in old queens) of sixty in a minute, or eighty
thousand and upward in one day of twenty-four hours."

This observer, after giving an account of the great swarms of perfect


winged Insects that are produced by this species, and after describing the
avidity with which they are devoured by the Hymenopterous ants and
other creatures, adds: "I have discoursed with several gentlemen upon
the taste of the white ants; and on comparing notes we have always
agreed that they are most delicious and delicate eating. One gentleman
compared them to sugared marrow, another to sugared cream and a
paste of sweet almonds."

From the preceding brief sketch of some Termitidae we may gather the
chief points of importance in which they differ from other Insects, viz. (1)
the existence in the community of individuals—workers and soldiers—
which do not resemble their parents; (2) the limitation of the reproductive
power to a single pair, or to a small number of individuals in each
community, and the prolongation of the terminal period of life. There are
other social Insects besides Termitidae: indeed, the majority of social
Insects—ants, bees, and wasps—belong to the Order Hymenoptera, and
it is interesting to note that analogous phenomena occur in them, but
nevertheless with such great differences that the social life of Termites
must be considered as totally distinct from that of the true ants and other
social Hymenoptera.

Development.—Social Insects are very different to others not only in the


fact of their living in society, but in respect of peculiarities in the mode of
reproduction, and in the variety of habits displayed by members of a
community. The greatest confusion has arisen in reference to Termitidae
in consequence of the phenomena of their lives having been assumed to
be similar to those of Hymenoptera; but the two cases are very different,
Hymenoptera passing the early parts of their lives as helpless maggots,
and then undergoing a sudden metamorphosis to a totally changed
condition of structure, intelligence, and instinct.

The development of what we may look on as the normal form of


Termitidae—that is, the winged Insects male and female—is on the whole
similar to that we have sketched in Orthoptera; the development in
earwigs being perhaps the most similar. The individuals of Termitidae are,
however, in the majority of cases if not in all, born without eyes; the wing-
rudiments develop from the thoracic terga as shorter or longer lobes
according to the degree of maturity; as in the earwigs the number of joints
in the antennae increases as development advances. All the young are,
when hatched, alike, the differences of caste appearing in the course of
the subsequent development; the most important of these differences are
those that result in the production of two special classes—only met with in
social Insects—viz. worker and soldier. Of these the workers are
individuals whose development is arrested, the sexual organs not going
on to their full development, while other organs, such as the eyes, also
remain undeveloped; the alimentary canal and its adjuncts occupy nearly
the whole of the abdominal cavity. The adult worker greatly resembles—
except in size—the young. Grassi considers that the worker is not a case
of simple arrest of development, but that some deviation accompanies the
arrest.

The soldier also suffers an arrest of development in certain respects


similar to the worker; but the soldier differs in the important fact that the
arrest of the development of certain parts is correlative with an
extraordinary development of the head, which ultimately differs greatly
from those of either the worker or of the sexual males and females.
Fig. 233.—The pairs of mandibles of different adult individuals of Termes sp.
from Singapore. A, Of worker; B, of soldier; C, of winged male; D, of
winged female.

Soldier.—All the parts of the head of the soldier undergo a greater or less
change of form; even the pieces at its base, which connect it by means of
the cervical sclerites with the prothorax, are altered. The parts that
undergo the greatest modification are the mandibles (Fig. 233, B); these
become much enlarged in size and so much changed in form that in a
great many species no resemblance to the original shape of these organs
can be traced. It is a curious fact that the specific characters are better
expressed in these superinduced modifications than they are in any other
part of the organisation (except, perhaps, the wings). The soldiers are not
alike in any two species of Termitidae so far as we know, and it seems
impossible to ascribe the differences that exist between the soldiers of
different species of Termitidae to special adaption for the work they have
to perform. Such a suggestion is justifiable only in the case of the Nasuti
(Fig. 234, 1), where the front of the head is prolonged into a point: a duct
opens at the extremity of this point, from which is exuded a fluid that
serves as a cement for constructing the nest, and is perhaps also used to
disable enemies. Hence the prolongation and form of the head of these
Nasuti may be fairly described as adaptation to useful ends. As regards
the great variety exhibited by other soldiers—and their variety is much
greater than it is in the Nasuti—it seems at present impossible to treat it
as being cases of special adaptations for useful purposes. On the whole it
would be more correct to say that the soldiers are very dissimilar in spite
of their having to perform similar work, than to state that they are
dissimilar in conformity with the different tasks they carry on.
Fig. 234.—Soldiers of different species of Termites. (After Hagen.) 1,
Termes armiger; 2, T. dirus; 3, Calotermes flavicollis; 4, T. bellicosus;
5, T. occidentis; 6, T. cingulatus (?); 7, Hodotermes quadricollis (?); 8,
T. debilis (?), Brazil.

The Termite soldier is a phenomenon to which it is difficult to find a


parallel among Insects. The soldier in the true ants is usually not definitely
distinguished from the worker, but it is possible that in the leaf-cutting
ants, the so-called soldier may prove to be more similar in its nature to
the Termite soldier. The soldiers of any one species of Termite are
apparently extremely similar to one another, and there are no
intermediates between them and the other forms, except in the stages of
differentiation. But we must recollect that but little is yet known of the full
history of any Termite community, and it is possible that soldiers which in
the stage of differentiation promise to be unsatisfactory may be killed and
eaten,—indeed there is some evidence to this effect. There is too in
certain cases some difference—larger or smaller size being the most
important—between the soldiers of one species, which may possibly be
due to the different stage of development at which their differentiation
commenced.

It would at present appear that, notwithstanding the remarkable difference


in structure of the soldiers and workers of the white ants, there is not a
corresponding difference of instinct. It is true that soldiers do more of
certain things than workers do, and less of others, but this appears to be
due solely to their possession of such very different structures; and we
are repeatedly informed by Grassi that all the individuals in a community
take part, so far as they are able, in any work that is going on, and we find
also in the works of other writers accounts of soldiers performing acts that
one would not have expected from them. The soldiers are not such
effective combatants as the workers are. Dudley and Beaumont indeed
describe the soldiers as merely looking on while the workers fight.[295] So
that we are entitled to conclude that the actions of the soldiers, in so far
as they differ from those of the rest of the community, do so because of
the different organisation and structures of these individuals. We shall,
when speaking of food, point out that the condition of the soldier in
relation to food and hunger is probably different from that of the other
forms.

Various Forms of a Community.—The soldiers and workers are not the


only anomalous forms found in Termitid communities; indeed on
examining a large nest a variety of forms may be found that is almost
bewildering. Tables have been drawn up by Grassi and others showing
that as many as fifteen kinds may be found, and most of them may under
certain circumstances coexist. Such tables do not represent the results of
actual examination in any one case, and they by no means adequately
represent the number that, according to the most recent observations of
Grassi, may be present; but we give one taken from Grassi, as it conveys
some idea of the numerous forms that exist in certain communities. In this
table the arrangement, according to A, B, C, D, E, represents successive
stages of the development:—

Forms of Termes lucifugus. (After Grassi.) Zool. Anz. xii. 1889, p. 360.
On inspecting this table it will be perceived that the variety of forms is due
to three circumstances—(1) the existence of castes that are not present
in ordinary Insects; (2) the coexistence of young, of adolescents, and of
adults; and (3) the habit the Termites have of tampering with forms in their
intermediate stages, the result of which may be the substitution of
neoteinic individuals in place of winged forms.

This latter procedure is far from being completely understood, but to it are
probably due the various abnormal forms, such as soldiers with rudiments
of wings, that have from time to time been discovered in Termite
communities, and have given rise to much perplexity.

In connexion with this subject we may call attention to the necessity, when
examining Termite nests, of taking cognisance of the fact that more than
one species may be present. Bates found different Termites living
together in the Amazons Valley, and Mr. Haviland has found as many as
five species of Termitidae and three of true ants in a single mound in
South Africa. In this latter case observation showed that, though in such
close proximity, there was but little further intimacy between the species.
There are, however, true inquiline, or guest, Termites, of the genus
Eutermes, found in various parts of the world living in the nests of other
Termitidae.

Origin of the Forms.—The interest attaching to the various forms that


exist in Termites, more particularly to the worker and soldier, is evident
when we recollect that these never, so far as we know, produce young. In
the social Hymenoptera it has been ascertained that the so-called neuters
(which in these Insects are always females) can, and occasionally do,
produce young, but in the case of the Termites it has never been
suggested that the sexual organs of the workers and soldiers, whether
male or female, ever become fruitful; moreover, the phenomena of the
production of young by the white ants are of such a nature as to render it
in the highest degree improbable that either workers or soldiers ever take
any direct part in it. Now the soldier is extremely different from the sexual
individuals that produce the young, and seeing that its peculiarities are
not, in the ordinary sense of the word, hereditary, it must be of great
interest to ascertain how they arise.
Before stating the little information we possess on this subject, it is
necessary to reiterate what we have already said to the effect that the
soldiers and workers are not special to either sex, and that all the young
are born alike. It would be very natural to interpret the phenomena by
supposing the workers to be females arrested in their development—as is
the case in social Hymenoptera—and by supposing the soldiers to be
males with arrested and diverted development.

The observations already made show that this is not the case. It has been
thoroughly well ascertained by Lespès and Fritz Müller that in various
species of Calotermes the soldiers are both males and females. Lespès
and Grassi have shown that the workers of Termes lucifugus are of male
and female sex, and that this is also true of the soldiers. Although the
view of the duality of the sexes of these forms was received at first with
incredulity, it appears to be beyond doubt correct. Grassi adds that in all
the individuals of the workers and soldiers of Termes lucifugus the sexual
organs, either male or female, are present, and that they are in the same
stage of development whatever the age of the individual. This statement
of Grassi's is of importance because it seems to render improbable the
view that the difference of form of the soldier and worker arises from the
arrest of the development of their sexual organs at different periods. The
fact that sex has nothing whatever to do with the determination of the
form of workers and soldiers may be considered to be well established.

The statement that the young are all born alike is much more difficult to
substantiate. Bates said that the various forms could be detected in the
new-born. His statement was made, however, merely from inspection of
the nests of species about which nothing was previously known, and as it
is then very difficult to decide that a specimen is newly hatched, it is
probable that all he meant was that the distinction of workers, soldiers,
and sexual forms existed in very small individuals—a statement that is no
doubt correct. Other observers agree that the young are in appearance all
alike when hatched, and Grassi reiterates his statement to this effect.
Hence it would appear that the difference of form we are discussing
arises from some treatment subsequent to hatching. It may be suggested,
notwithstanding the fact that the young are apparently alike when
hatched, that they are not really so, but that there are recondite
differences which are in the course of development rendered
conspicuous. This conclusion cannot at present be said with certainty to
be out of the question, but it is rendered highly improbable by the fact
ascertained by Grassi that a specimen that is already far advanced on the
road to being an ordinary winged individual can be diverted from its
evident destination and made into a soldier, the wings that were partially
developed in such a case being afterwards more or less completely
absorbed. This, as well as other facts observed by Grassi, render it
probable that the young are truly, as well as apparently, born in a state
undifferentiated except as regards sex. Fig. 230 (p. 363) is designed to
illustrate Grassi's view as to this modification; the individual A is already
far advanced in the direction of the winged form C, but can nevertheless
be diverted by the Termites to form the adult soldier B.

According to the facts we have stated, neither heredity nor sex nor arrest
of development are the causes of the distinctions between worker and
soldier, though some arrest of development is common to both; we are
therefore obliged to attribute the distinction between them to other
influences. Grassi has no hesitation in attributing the anatomical
distinctions that arise between the soldiers, workers, and winged forms to
alimentation. Food, or the mode of feeding, or both combined, are,
according to the Italian naturalist, the source of all the distinctions, except
those of sex, that we see in the forms of any one species of Termite.

Feeding.—Such knowledge as we possess of the food-habits of


Termitidae is chiefly due to Grassi; it is of the very greatest importance, as
giving a clue to much that was previously obscure in the Natural History
of these extraordinary creatures.

In the abodes of the Termites, notwithstanding the enormous numbers of


individuals, cleanliness prevails; the mode by which it is attained appears
to be that of eating all refuse matter. Hence the alimentary canal in
Termitidae contains material of various conditions of nutritiveness. These
Insects eat their cast skins and the dead bodies of individuals of the
community; even the material that has passed through the alimentary
canal is eaten again, until, as we may presume, it has no further nutritive
power. The matter is then used for the construction of their habitations or
galleries, or is carried to some unfrequented part of the nest, or is voided
by the workers outside of the nests; the pellets of frass, i.e. alimentary
rejectamenta, formed by the workers frequently betraying their presence
in buildings when none of the Insects themselves are to be seen. The
aliments of Calotermes flavicollis are stated by Grassi and Sandias to be
as follows: (1) wood; (2) material passed from the posterior part of the
alimentary canal or regurgitated from the anterior part; (3) the matter shed
during the moults; (4) the bodies of other individuals; (5) the secretion of
their own salivary glands or that of their fellows; (6) water. Of these the
favourite food is the matter passed from the posterior part of the
alimentary canal. We will speak of this as proctodaeal food. When a
Calotermes wishes food it strokes the posterior part of another individual
with the antennae and palpi, and the creature thus solicited yields, if it
can, some proctodaeal food, which is then devoured. Yielding the
proctodaeal food is apparently a reflex action, as it can be induced by
friction and slight pressure of the abdomen with a small brush. The
material yielded by the anterior part of the alimentary canal may be called
stomodaeal product. It makes its appearance in the mouth in the form of a
microscopic globule that goes on increasing in size till about one
millimetre in diameter, when it is either used for building or as food for
another individual. The mode of eating the ecdysial products has also
been described by Grassi and Sandias. When an individual is sick or
disabled it is frequently eaten alive. It would appear that the soldiers are
great agents in this latter event, and it should be noticed that owing to
their great heads and mandibles they can obtain food by other means
only with difficulty. Since they are scarcely able to gnaw wood, or to
obtain the proctodaeal and stomodaeal foods, their condition may be
considered to be that of permanent hunger, only to be allayed by
carnivorous proceedings. When thrown into a condition of excitement the
soldiers sometimes exhibit a sort of Calotermiticidal mania, destroying
with a few strokes five or six of their fellows. It is, however, only proper to
say that these strokes are made at random, the creature having no eyes.
The carnivorous propensities of Calotermes are apparently limited to
cannibalism, as they slaughter other white ants (Termes lucifugus) but
never eat them.

The salivary food is white and of alkaline nature; when excreted it makes
its appearance on the upper lip. It is used either by other individuals or by
the specimen that produced it; in the latter case it is transferred to the
lower lip and swallowed by several visible efforts of deglutition. The
aliments we have mentioned are made use of to a greater or less extent
by all the individuals except the very young; these are nourished only by
saliva: they commence taking proctodaeal and stomodaeal food before
they can eat triturated wood.

Royal Pairs.—The restriction of the reproductive powers of a community


to a single pair (or to a very restricted number of individuals) occurs in all
the forms of social Insects, and in all of them it is concomitant with a
prolongation of the reproductive period far exceeding what is natural in
Insects. We are not in a position at present to say to what extent the lives
of the fertile females of Termitidae are prolonged, there being great
difficulties in the way of observing these Insects for long periods owing to
their mode of life; living, as they do, concealed from view, light and
disturbance appear to be prejudicial to them. We have every reason to
believe, however, that the prolongation extends as a rule over several
years, and that it is much greater than that of the other individuals of the
community, although the lives of even these latter are longer than is usual
in Insects; but this point is not yet satisfactorily ascertained. As regards
the males there is reason to think that considerable variety as to longevity
prevails. But the belief is that the royal males of Termitidae also form an
exception to other Insects in the prolongation of the terminal periods of
their lives. In Hymenoptera, male individuals are profusely produced, but
their lives are short, and their sole duty is the continuation of the species
by a single act. We have seen that Grassi is of opinion that a similar
condition of affairs exists at present with Termes lucifugus in Sicily, but
with this exception it has always been considered that the life of the king
Termite is, roughly speaking, contemporaneous with that of the queen; it
is said that in certain species the king increases in bulk, though not to an
extent that can be at all compared with the queen.

It must be admitted that the duration of life of the king has not been
sufficiently established, for the coexistence of a king with a queen in the
royal cell is not inconsistent with the life of the king being short, and with
his replacement by another. Much that is imaginary exists in the literature
respecting Termites, and it is possible that the life of the king may prove
to be not so prolonged as has been assumed.
Fig. 235.—Royal pair of Termes sp. from Singapore, taken out of royal cell.
A, A, King, lateral and dorsal views; B, B, queen, dorsal and lateral
views. Natural sizes.

Returning to the subject of the limitation of the reproduction of the


community to a single pair, we may remark that a priori one would
suppose such a limitation to be excessively unfavourable to the
continuation of the species; and as it nevertheless is the fact that this
feature is almost, if not quite, without exception in Insect societies, we
may conclude that it is for some reason absolutely essential to Insect
social life. It is true that there are in Termitidae certain partial exceptions,
and these are so interesting that we may briefly note them. When a royal
cell is opened it usually contains but a single female and male, and when
a community in which royal cells are not used is inspected it is usually
found that here also there are present only a single fertile female and a
single king. Occasionally, however, it happens that numerous females are
present, and it has been noticed that in such cases they are not fully
matured females, but are imperfect, the condition of the wings and the
form of the anterior parts of the body being that of adolescent, not adult
Insects. It will be recollected that the activity of a community of Termites
centres round the great fertility of the female; without her the whole
community is, as Grassi graphically puts it, orphaned; and the
observations of the Italian naturalist make it clear that these imperfect
royalties are substitution queens, derived from specimens that have not
undergone the natural development, but have been brought into use to
meet the calamity of orphanage of the community. The Termites
apparently have the power of either checking or stimulating the
reproductive organs apart from other organs of the body, and they appear
to keep a certain number of individuals in such a condition that in case of
anything going wrong with the queen, the reserves may be brought as
soon as possible into a state of reproductive activity. The individuals that
are in such a condition that they can become pseudo-royalties are called
complementary or reserve royalties, and when actually brought into use
they become substitution royalties. It is not at present quite clear why the
substitution royalties should be in such excess of numbers as we have
stated they were in the case we have figured (Fig. 236), but it may be due
to the fact that when the power of the community is at a certain capacity
for supporting young a single substitution royalty would not supply the
requisite producing power, and consequently the community adopts a
greater number of the substitution forms. Termites are utterly regardless
of the individual lives of the members of the community, and when the
reproductive powers of the company of substitution royalties become too
great, then their number is reduced by the effective method of killing and
eating them.

According to Grassi's observations, the communities of Termes lucifugus


are now kept up in Sicily almost entirely by substitution royalties; the
inference being that the age of each community has gone beyond the
capacity for life of any single royal queen.

The substitution royalties are, as we have said, called neoteinic (νεος,


youthful, τείνω, to belong to), because, though they carry on the functions
of adult Insects, they retain the juvenile condition in certain respects, and
ultimately die without having completed the normal development. The
phenomenon is not quite peculiar to Insects, but occurs in some other
animals having a well-marked metamorphosis, notably in the Mexican
Axolotl.[296]

Fig. 236.—Pair of neoteinic royalties, taken from the royal chamber of


Termes sp. at Singapore by Mr. G. D. Haviland. The queen was one of
thirteen, all in a nearly similar state. A, king; B, C, queen.

A point of great importance in connexion with the neoteinic royalties is


that they are not obtained from the instar immediately preceding the adult
state, but are made from Insects in an earlier stage of development. The
condition immediately preceding the adult state is that of a nymph with
long wing-pads; such specimens are not made into neoteinic royalties,
but nymphs of an earlier stage, or even larvae, are preferred. It is
apparently by an interference with one of these earlier stages of
development that the "nymphs of the second form," which have for long
been an enigma to zoologists, are produced.

Post-metamorphic Growth.—The increase of the fertility of the royal


female is accompanied by remarkable phenomena of growth. Post-
metamorphic growth is a phenomenon almost unknown in Insect life,
except in these Termitidae; distension not infrequently occurs to a certain
extent in other Insects, and is usually due to the growth of eggs inside the
body, or to the repletion of other parts. But in Termitidae there exists post-
metamorphic growth of an extensive and complex nature; this growth
does not affect the sclerites (i.e. the hard chitinous parts of the exo-
skeleton), which remain of the size they were when the post-metamorphic
growth commenced, and are consequently mere islands in the distended
abdomen (Fig. 236, B, C). The growth is chiefly due to a great increase in
number and size of the egg-tubes, but there is believed to be a correlative
increase of various other parts of the abdominal as distinguished from the
anterior regions of the body. A sketch of the distinctions existing between
a female of a species at the time of completion of the metamorphosis and
at the period of maximum fertility does not appear to have been yet made.

New Communities.—The progress of knowledge in respect of Termitidae


is bringing to light a quite unexpected diversity of habits and constitution.
Hence it is premature to generalise on important matters, but we may
refer to certain points that have been ascertained in connexion with the
formation of new communities. The duration of particular communities
and the modes in which new ones are founded are still very obscure. It
was formerly considered that swarming took place in order to increase the
number of communities, and likewise for promoting crossing between the
individuals of different communities. Grassi, however, finds as the result
of his prolonged observations on Termes lucifugus that the swarms have
no further result than that the individuals composing them are eaten up.
And Fritz Müller states[297] that in the case of the great majority of forms
known to him the founding of a colony by means of a pair from a swarm
would be just about as practicable as to establish a new colony of human
beings by placing a couple of newly-born babes on an uninhabited island.
It was also thought that pairs, after swarming, re-entered the nests and
became royal couples. It does not, however, appear that any one is able
to produce evidence of such an occurrence. The account given by
Smeathman of the election of a royal couple of Termes bellicosus is
imperfect, as, indeed, has already been pointed out by Hagen. It
suggests, however, that a winged pair after leaving the nest do again
enter it to become king and queen. The huge edifices of this species
described by Smeathman are clearly the result of many years of labour,
and at present substitution royalties are not known to occur in them, so
that it is not improbable Smeathman may prove to be correct even on this
point, and that in the case of some species mature individuals may re-
enter the nest after swarming and may become royal couples. On the
whole, however, it appears probable that communities of long standing
are kept up by the substitution royalty system, and that new communities
when established are usually founded by a pair from a swarm, which at
first are not in that completely helpless condition to which they come
when they afterwards reach the state of so-called royalty. Grassi's
observations as to the sources of food remove in fact one of the
difficulties that existed previously in regard to the founding of new
colonies, for we now know that a couple may possibly bear with them a
sufficient supply of proctodaeal and stomodaeal aliment to last them till
workers are hatched to feed them, and till soldiers are developed and the
community gradually assumes a complex condition. Professor Perez has
recently obtained[298] the early stages of a community from a winged pair
after they had been placed in captivity, unattended by workers. Müller's
observation, previously quoted, is no doubt correct in relation to the
complete helplessness of royal pairs after they have been such for some
time; but that helplessness is itself only gradually acquired by the royal
pair, who at first are able to shift for themselves, and produce a few
workers without any assistance.

Anomalous Forms.—Müller has described a Calotermes under the


name of C. rugosus, which is interesting on account of the peculiar form
of the young larva, and of the changes by which it subsequently becomes

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